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      Teaching varies with task complexity in wild chimpanzees

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          Significance

          Understanding social influences on how apes acquire tool behaviors can help us model the evolution of culture and technology in humans. Humans scaffold novice tool skills with diverse strategies, including the transfer of tools between individuals. Chimpanzees transfer tools, and this behavior meets criteria for teaching. However, it is unclear how task complexity relates to this form of helping. Here, we find differences between 2 wild chimpanzee populations in rate, probability, and types of tool transfer during termite gathering. Chimpanzees showed greater helping in the population where termite gathering is a more complex tool task. In wild chimpanzees, as in humans, regular and active provisioning of learning opportunities may be essential to the cultural transmission of complex skills.

          Abstract

          Cumulative culture is a transformative force in human evolution, but the social underpinnings of this capacity are debated. Identifying social influences on how chimpanzees acquire tool tasks of differing complexity may help illuminate the evolutionary origins of technology in our own lineage. Humans routinely transfer tools to novices to scaffold their skill development. While tool transfers occur in wild chimpanzees and fulfill criteria for teaching, it is unknown whether this form of helping varies between populations and across tasks. Applying standardized methods, we compared tool transfers during termite gathering by chimpanzees in the Goualougo Triangle, Republic of Congo, and in Gombe, Tanzania. At Goualougo, chimpanzees use multiple, different tool types sequentially, choose specific raw materials, and perform modifications that improve tool efficiency, which could make it challenging for novices to manufacture suitable tools. Termite gathering at Gombe involves a single tool type, fishing probes, which can be manufactured from various materials. Multiple measures indicated population differences in tool-transfer behavior. The rate of transfers and probability of transfer upon request were significantly higher at Goualougo, while resistance to transfers was significantly higher at Gombe. Active transfers of tools in which possessors moved to facilitate possession change upon request occurred only at Goualougo, where they were the most common transfer type. At Gombe, tool requests were typically refused. We suggest that these population differences in tool-transfer behavior may relate to task complexity and that active helping plays an enhanced role in the cultural transmission of complex technology in wild apes.

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          Ratcheting up the ratchet: on the evolution of cumulative culture.

          Some researchers have claimed that chimpanzee and human culture rest on homologous cognitive and learning mechanisms. While clearly there are some homologous mechanisms, we argue here that there are some different mechanisms at work as well. Chimpanzee cultural traditions represent behavioural biases of different populations, all within the species' existing cognitive repertoire (what we call the 'zone of latent solutions') that are generated by founder effects, individual learning and mostly product-oriented (rather than process-oriented) copying. Human culture, in contrast, has the distinctive characteristic that it accumulates modifications over time (what we call the 'ratchet effect'). This difference results from the facts that (i) human social learning is more oriented towards process than product and (ii) unique forms of human cooperation lead to active teaching, social motivations for conformity and normative sanctions against non-conformity. Together, these unique processes of social learning and cooperation lead to humans' unique form of cumulative cultural evolution.
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            Hunting behavior of wild chimpanzees in the Taï National Park.

            Hunting is often considered one of the major behaviors that shaped early hominids' evolution, along with the shift toward a drier and more open habitat. We suggest that a precise comparison of the hunting behavior of a species closely related to man might help us understand which aspects of hunting could be affected by environmental conditions. The hunting behavior of wild chimpanzees is discussed, and new observations on a population living in the tropical rain forest of the Taï National Park, Ivory Coast, are presented. Some of the forest chimpanzees' hunting performances are similar to those of savanna-woodlands populations; others are different. Forest chimpanzees have a more specialized prey image, intentionally search for more adult prey, and hunt in larger groups and with a more elaborate cooperative level than savanna-woodlands chimpanzees. In addition, forest chimpanzees tend to share meat more actively and more frequently. These findings are related to some theories on aspects of hunting behavior in early hominids and discussed in order to understand some factors influencing the hunting behavior of wild chimpanzees. Finally, the hunting behavior of primates is compared with that of social carnivores.
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              Identification of the social and cognitive processes underlying human cumulative culture.

              The remarkable ecological and demographic success of humanity is largely attributed to our capacity for cumulative culture, with knowledge and technology accumulating over time, yet the social and cognitive capabilities that have enabled cumulative culture remain unclear. In a comparative study of sequential problem solving, we provided groups of capuchin monkeys, chimpanzees, and children with an experimental puzzlebox that could be solved in three stages to retrieve rewards of increasing desirability. The success of the children, but not of the chimpanzees or capuchins, in reaching higher-level solutions was strongly associated with a package of sociocognitive processes-including teaching through verbal instruction, imitation, and prosociality-that were observed only in the children and covaried with performance.
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                Author and article information

                Journal
                Proc Natl Acad Sci U S A
                Proc. Natl. Acad. Sci. U.S.A
                pnas
                pnas
                PNAS
                Proceedings of the National Academy of Sciences of the United States of America
                National Academy of Sciences
                0027-8424
                1091-6490
                14 January 2020
                23 December 2019
                23 December 2019
                : 117
                : 2
                : 969-976
                Affiliations
                [1] aDepartment of Anthropology, University of Miami , Coral Gables, FL 33124;
                [2] bDepartment of Psychology, Franklin and Marshall College , Lancaster, PA 17604;
                [3] cBiological Foundations of Behavior Program, Franklin and Marshall College , Lancaster, PA 17604;
                [4] dLester E. Fisher Center for the Study and Conservation of Apes, Lincoln Park Zoo , Chicago, IL 60614;
                [5] eDepartment of Biology, Case Western Reserve University , Cleveland, OH 44106;
                [6] fConservation and Science Department, Cleveland Metroparks Zoo , Cleveland, OH 44109;
                [7] gMax Planck Institute for Evolutionary Anthropology , 04103 Leipzig, Germany;
                [8] hDepartment of Anthropology, Washington University in St. Louis , St. Louis, MO 63130;
                [9] iCongo Program, Wildlife Conservation Society , B.P. 14537 Brazzaville, Republic of Congo;
                [10] jKyoto University Institute for Advanced Study, Kyoto University , Kyoto, 606-8501, Japan
                Author notes
                1To whom correspondence may be addressed. Email: smusgrave@ 123456miami.edu .

                Edited by Frans B. M. de Waal, Emory University, Atlanta, GA, and approved November 12, 2019 (received for review May 7, 2019)

                Author contributions: S.M., E.L., D.M., and C.S. designed research; S.M., E.L., D.M., M.P., L.B.-K., and C.S. performed research; R.M. analyzed data; and S.M., D.M., and C.S. wrote the paper.

                Article
                201907476
                10.1073/pnas.1907476116
                6969499
                31871167
                01565593-647c-41e2-827c-829463236c38
                Copyright © 2020 the Author(s). Published by PNAS.

                This open access article is distributed under Creative Commons Attribution-NonCommercial-NoDerivatives License 4.0 (CC BY-NC-ND).

                History
                Page count
                Pages: 8
                Categories
                PNAS Plus
                Biological Sciences
                Anthropology
                From the Cover
                PNAS Plus

                tool use,cumulative culture,social learning,chimpanzee,prosociality

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